ACTA PALAEONTOLOGICA ROMANIAE (2018) V. 14 (1), P. 31-45

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1Geological Institute of Romania - 1st, Caransebeş Street, Ro-012271, Bucharest, Romania. E-mail: iamandei@gmail.com 31 2Istanbul University Forestry Faculty, Department of Forest Botany, 34473 Bahceköy-Sarıyer, İstanbul / Turkey.

*Corresponding author: uakkemik@istanbul.edu.tr

NEOGENE PALMOXYLON FROM TURKEY

Stănilă Iamandei1, Eugenia Iamandei1 & Ünal Akkemik2*

Received: 13 May 2018 / Accepted: 26 July 2018 / Published online: 2 August 2018

Abstract Tertiary fossil wood identifications of Turkey revealed many different gymnosperm and angiosperm genera

and species. Within this lignoflora some Palmoxylon types were identified for the first time as Palmoxylon coryph-

oides Ambwani et Mehrotra, Palmoxylon sp. cf. Trachycarpus H. Wendl., and Palmoxylon sp. cf. Borassus L. The

fossil palm-wood was collected from the sites in Seben and Gökçeada from early Miocene deposits, and from the site

in Erikli from middle-late Miocene deposits. The presence of the palm trees in the Miocene Flora of the Anatolian

and Aegean regions indicates a lowland riparian palaeoenvironment and warm-humid or subtropical palaeoclimate.

Keywords: Palmoxylon, fossil-species, Seben, Gökçeada, Erikli.

INTRODUCTION

In the recent decades, our information on the fossil lig-

noflora of Turkey has increased. The new identifications

of petrified wood in Turkey showed a rich woody flora

from the late Oligocene to the late Miocene. Some of the

identified genera such as Sequoioxylon (Özgüven-Ertan,

1971, Kayacik et al., 1995; Akkemik et al., 2005; Sakınç

et al., 2007, Akkemik & Sakinç, 2013, Akkemik et al.,

2009), Glyptostroboxlon (Akkemik et al., 2017), Podo-

carpoxylon (Sakınç et al., 2007), Engelhardioxylon

(Sakınç et al., 2007) are taxa completely extinct from

Turkey, while other fossil forms (e.g. Sayadi, 1973; Sel-

meier, 1990; Dernbach et al., 1996; Aytuğ & Şanli, 1974;

Eroskay & Aytug, 1982; Şanli, 1982; Akkemik et al.,

2016 and Bayam et al., 2018) still have modern corre-

spondents. Regarding with Palmoxylon, many studies

were published in the world (e.g. Schenk, 1882; Mahaba-

lé, 1958; Greguss, 1959, 1969; Kaul, 1960; Tomlinson,

1961, 1990; Prakash, 1962; Grambast, 1962, 1964;

Trivedi & Verma, 1970; Roy & Ghosh, 1980; Privé-Gill

& Pelletier, 1981; Ambwani & Mehrotra, 1990; Gott-

wald, 1992; Nambudiri & Tidwell, 1998; EL-Saadawi et

al., 2004; Kahlert et al., 2005; Sakala, 2004; Dransfield et

al., 2008; Thomas, 2011a,b; Thomas & De Franceschi,

2012, 2013; Kamal-EL-Din et al., 2013; Prasad et al.,

2013; Nour-EL-Deen et al., 2017). Otherwise an extend-

ed list o References on Palmoxylon is published in the

site CiteULike by Millevacs in: (Accessed 05.05.2018).

Within these studies Kamal-EL-Din et al. (2013) stated

that Egypt is the richest African country having fossil-

species of Palmoxylon with 16 species, after describing

four Palmoxylon species, P. deccanense Sahni, P. ed-

wardsi Sahni, P. geometricum Sahni, P. prismaticum

Sahni, P. pondicherriense Sahni, P. pyriforme Sahni and

P. sagari Sahni.

Thomas & De Franceschi (2013) made a valuable revi-

sion of the palm xylotomy, which is very useful for mod-

ern palm identification by the study of the palm stem mi-

croscopic structure and also useful for the fossil palms, as

species of Palmoxylon. Descriptions were based mainly

the general organization of the transverse section and the

structure of the fibrovascular bundles as well as the

ground parenchyma, taking the Coryphoideae as case-

study.

As for Anatolian region, the palaeobotanical studies men-

tioned above revealed that the climate in the early Mio-

cene was warmer, probably subtropical and much humid

than the present climate. Palm trees growth even today

under this kind of climate in Seben (Akkemik et al.,

2016), European Part of Turkey and in the Island of

Gökçeada in Aegean Sea (Güngör et al., 2018), but their

distribution area was much wider through Turkey during

Neogene. Under this discussion, the purpose of the pre-

sent paper is to describe some fossil forms based on the

study of their petrified remains.

MATERIAL AND METHODS

We had in study four samples that sowed the typical palm

structure and were considered as fossil forms of Palmoxy-

lon. They were coded as HOC25 (Akkemik et al., 2016),

GOK01 and GOK269 (Güngör et al., 2018), and ERI01.

They have been collected from Bolu-Seben-Hoçaş Fossil

Site (HOC25), Edirne-Erikli (ERI01) and Gökçeada

(GOK01 and GOK269) (Fig. 1). The samples were in-situ

position in Bolu-Seben-Hoçaş Fossil Site (Fig. 2) and

dispersed in the others (Fig. 3).

According to Akkemik et al. (2016), the age of fossil site

in Seben-Hoçaş was 18.2±0.8 from the basalt samples.

Similar ages were given by Keller et al. (1992), Toprak et

al. (1996), and Wilson et al. (1997) including the ages of

17.6-24.8 Ma for volcanic activity in Galatean Volcanic

Province. So, the age of Seben-Hoçaş Fossil Site is the

early Miocene.

Güngör et al. (2018) specified that the geological age of

Gökçeada as also early Miocene, bearing a similar wood

flora with Lesbos Island (Süss & Velitzelos, 1997).

The fossil wood material (ERI01) in Erikli was found on

the shore of Aegean Sea. Sakinç et al. (1999) explained

geological background of Erikli, and its age is most prob-

ably middle-late Miocene.

Stănilă Iamandei, Eugenia Iamandei & Ünal Akkemik

32

Fig. 1 The locations of the sampled materials.

Fig. 2 In-situ trunk of Palmoxylon in Seben-Hoçaş Fossil Site.

Fig. 3 Dispersed wood trunk fragments including one of Palmoxylon in Gökçeada Fossil Site.

Neogene Palmoxylon from Turkey

33

For wood identification, we realized thin sections from

the samples of fossil wood along the three standard

planes: transverse section (TS), or cross-section, radial

longitudinal section (RLS) or simply, radial section and

tangential longitudinal section (TLS) or simply tangential

section. These standard planes of cutting in a palm stem

are also valid since there is an oriented structure in the

external and intermediary part, only in the central part of

the stem is less important (see Mahabalé, 1958; Kaul,

1960; Greguss, 1959, 1968; Tomlinson, 1961)

The samples and sections were stored at the "Laboratory

of Tree-ring Researches and Wood Anatomy" in Forest

Botany Department of Faculty of Forestry, Istanbul Uni-

versity.

The first study was made on a biological microscope

(LEICA DM2500 Light Microscope) and the micro-

photos were realized with a LEICA DSC295 type of

camera.

The description and identification of the specimens was

performed in the Geological Institute of Romania, in Bu-

charest. In identification, many references including fos-

sil Palmoxylon species were used (e.g. Schenk, 1882;

Mahabalé, 1958; Greguss, 1959, 1968, 1969; Kaul, 1960;

Tomlinson, 1961, 1990; Prakash, 1962; Grambast, 1962,

1964; Trivedi & Verma, 1970; Roy & Ghosh, 1980;

Privé-Gill & Pelletier, 1981; Ambwani & Mehrotra,

1990; Gottwald, 1992; Nambudiri & Tidwell, 1998; EL-

Saadawi et al., 2004; Kahlert et al., 2005; Sakala, 2004;

Dransfield et al., 2008; Thomas, 2011a, b; Thomas & De

Franceschi, 2012, 2013; Kamal-EL-Din et al., 2013; Pra-

sad et al. 2013; Nour-El-Deen et al., 2017).

SYSTEMATIC PALAEOBOTANY

Order Arecales Bromhead 1840 (in Reveal, 2004)

Family Arecaceae Berchtold et Presl, 1820 (nom. cons)

Genus Palmoxylon Schenk, 1882

Palmoxylon coryphoides Ambwani & Mehrotra, 1990

Fig. 4, photos a-i. Fig. 5, photos a-i.

Material: GOK01 and GOK259.

Locality: Gökçeada.

Formation: Kesmekaya Volcanics.

Age: Early Miocene.

Origin on the stem remains: Coming from unknown

portion of the trunk.

Storage: This material is now kept under index GOK01

and GOK269 within the "Collection of fossil woods" in

the Forest Botany Department, Forestry Faculty, Istanbul

University.

Microscopic description: Theoretically the central cyl-

inder, in cross section, is formed from three distinct parts:

dermal, subdermal and central zones (see Mahabalé, 1958

and Kaul, 1960 quoting von Mohl, 1850) or external,

intermediary and central (in euro-american school, see

Tomlinson, 1961), or, respectively, subcortical zone - as a

periferal sclerotic zone, transitional zone and central zone

as was lastly stated by Thomas & De Franceschi (2013).

For description, the shape, the size, the density of the

fibrovascular bundles, their fibrous part and vessels, the

quantity of parenchyma, the abundance and the arrange-

ment of parenchymatous cells starch grained along to the

fibrovascular bundles must be considered and also the

phytoliths.

The studied specimens show in cross-section the central

zone, but also at least a part of the transitional zone (es-

pecially the specimen GOK259), where fibro-vascular

bundles of open collateral type appear, usually oriented

with the vascular part to the centre of the stem and the

fibrous part to outside, and having 2-4 large metaxylem

vessels round to oval, and typical reniform sclerenchyma

caps well developed, floating in the ground parenchyma

tissue, often touching each other. The size of fvb is varia-

ble 400-500/360-475 µm (mean values: 421.25 / 403.75

μm, the phloematic (=anterior) sclerenchyma cap has

r/tg.d = 116 / 372 μm, f / v ratio is 0.35 / 1 and the fre-

quency, or density, is of 491.2 bundles/cm2.

The phloem appears protected under the median sinus of

the sclerenchyma cap, a single island - as an undivided

sieve plate, rarely apparently divided.

The metaxylem appear as 2-4(-6-8) vessels either oval or

elliptic or, less round in cross-section or, sometimes, par-

titioned by the terminal inclined wall. They are usually

large and thick-walled vessels, having in cross section 70-

162.5 / 45-107.5 μm (mean values 107.7 / 65 μm) in di-

ameters and 8-10 µm the simple wall. The size of meta-

xylem vessels is of 47 / 44.55 μm, and thick-walled, of 7-

10 µm the simple wall. Vertically the walls of metaxylem

vessels have scalariform pitting and annular thickenings

sometimes ramified and anastomosed. The terminal wall

is very inclined and bears a scalariform perforation with

8-18 thick bars and, sometimes, only simple perforation.

The protoxylem appear as unequally smaller vessels

grouped beneath the metaxylem vessels or partially

pushed between them, possibly because the sustained

growing of the parenchyma from around. They have

d=25/22.9 μm, and are relatively thick-walled: 3-5 μm in

the simple wall.

The intrafascicular (or paravascular) parenchyma is con-

stituted of uniform, relatively thin-walled (5-7 μm the

simple wall), rounded-polyhedral cells which appear also

rounded in cross-section and small-sized (15.5/5 - 25/10

μm). Adjacent to the metaxylem vessels appear in 1-2

rows regularly arranged, similar to a tabular parenchyma.

The marginal 1-2 rows of sclerified paravascular paren-

chyma cells constitute a second sclerenchyma cap pro-

tecting the vascular zone also named ventral fibrous cap

adjacent to the xylem.

The ground tissue (or the interfascicular parenchyma) is

constituted by non-oriented rounded-polyhedral cells

unequally sized, very emaciated or lobed cells, as they

appear in cross section, usually thin-walled, and with

possible starch-content. It represents the ground paren-

chyma with sustained growing of Corypha type. All

around the fibrovascular bundle only tabular parenchyma

can be seen, in 1-2 compact rows arranged. Fibrous bun-

dles weren't observed.

The central zone in both the specimens studied is of Cor-

ypha type (von Mohl, in Thomas & De Franceschi, 2011)

where the round to oval fibrovascular bundles are varia-

bly oriented, having 2-4(-6-8) metaxylem vessels and not

too developed sclerenchyma caps of Reniforma to Lunar-

ia type, all floating in the ground parenchymal tissue

along with small fibrous bundles, more numerous in

Stănilă Iamandei, Eugenia Iamandei & Ünal Akkemik

34

Fig. 4 Palmoxylon coryphoides Ambwani & Mehrotra, 1990. (GOK01). a-c Cross-section. Fibrovascular bundles (fvb)

variably oriented in central zone; parenchymal ground tissue; fibrous bundles; c Detailed fvb-s displaying metaxylem and

protoxylem vessels, phloem, reniform sclerenchyma cap, stegmata, and rounded by tabular parenchyma. Also ground

tissue parenchymal cells and fibrous bundles can be seen. d-i Longitudinal sections. Metaxylem and protoxylem vessels

with annular thickenings, scalariform pitting, and with scalariform perforated inclined plates. Also, long rows of stegmata

can be seen. g, i Metaxylem and protoxylem vessels with annular bifurcated thickenings, and with scalariform perforated

plate. h Metaxylem vessel with scalariform pitting.

Neogene Palmoxylon from Turkey

35

Fig. 5 Palmoxylon coryphoides Ambwani & Mehrotra, 1990. (GOK259). a-c Cross-section. a Transitional zone with fvb

with reniform caps; parenchymal ground tissue; b Variably oriented fvb in central zone; parenchymal ground tissue; fi-

brous bundles; c Detailed fvb coupled with small fvb-s displaying metaxylem and protoxylem vessels, phloem, reniform

sclerenchyma cap, stegmata, and tabular parenchyma. Also ground tissue parenchymal cells and fibrous bundles can be

seen. d-g Longitudinal sections. Metaxylem and protoxylem vessels with annular or helical thickenings, scalariform pit-

ting, and with scalariform perforated inclined plates. Fibrous sheaths of fvb marked by long rows of stegmata (d, f). g-i

Metaxylem and protoxylem vessels with annular and helical thickenings and with simple perforation (i).

Stănilă Iamandei, Eugenia Iamandei & Ünal Akkemik

36

GOK01 specimen. Under the sclerenchyma cap a large

median sinus is present, auricular sinuses were not ob-

served. The sclerenchyma cap (also named anterior or

dorsal cap, or phloematic fibrous sheath, or fibrous part

adjacent to phloem) is constituted from vertically elon-

gated fibrous cells with polygonal rounded cross-section,

thick-walled and with rounded to point-like lumina. The

size of the fibrovascular bundles is variable: 400-

500/360-475 µm (mean values: 421.25/403.75 μm, the

phloematic (=anterior) sclerenchyma cap has

r/tg.d=116/372 μm, f/v ratio is 0.35/1 and the frequency

(or density) is of 214 bundles/cm2. Signs of a centrifugal

differentiation of the fibrous part seem to be absent.

The phloem, appear protected under the median sinus of

the sclerenchyma cap, in a single island - as an undivided

sieve plate, rarely apparently divided.

The metaxylem appear as 2-4(-6-8) vessels either oval or

elliptic, or less, round in cross-section or, sometimes par-

titioned by the terminal wall. They are usually large and

thick-walled vessels (7-8-10 µm the simple wall), having,

in cross section, 70-162.5/45-107.5 µm (mean 107.7/65

µm) in diameters (smaller in specimen GOK259:

47.03/44.55). Vertically the walls of the metaxylem ves-

sels have scalariform pitting and annular thickenings

sometimes ramified. The terminal wall is very inclined

and bears a scalariform perforation with 8-18 thick bars

(fewer in the specimen GOK259: 3-5-8 bars) or even

simple perforation.

The protoxylem is represented by 3-12 small round to

oval vessels of 10-36 µm in diameters, more numerous in

the specimen GOK259 (up to 30), and relatively thick

walled (2-5 µm the simple wall). Vertically the protoxy-

lem vessels present annular thickenings sometimes rami-

fied, and short scalariform perforations.

The intrafascicular (or paravascular) parenchyma is con-

stituted of uniform, thin-walled (1-2 µm the double wall),

rounded-polyhedral cells which appear polygonal-

rounded in cross-section, small-sized cells (round of 6-14

µm or elliptic of 12.5-25/5-15 µm). Adjacent to the meta-

xylem vessels appear in 1-2 regular rows of flattened

cells arranged, like a tabular parenchyma. All the re-

mained space is filled with compact non-oriented paren-

chyma cells. The marginal 1-3 rows of paravascular pa-

renchyma is constituted by highly sclerified cells, thicker

walled (3-6 µm the double wall), so forming a second

sclerenchyma cap, which protect the vascular zone, also

named ventral cap or fibrous part adjacent to the xylem.

The ground tissue or the interfascicular parenchyma is

constituted by non-oriented very emaciated big cells as

lobed cells, as they appear in cross section, usually thin-

walled, and with possible starch content. It represents the

ground parenchyma with sustained growing of Corypha

type, the cells compressing each other. All around the

fibrovascular bundle only tabular parenchyma can be

seen, in 1-2 compact rows arranged.

The fibrous bundles are more numerous in specimen

GOK01, floating in the ground parenchyma among the

fibrovascular bundles. They have 15-19-25 fibers, or

more, with polygonal rounded cross-section, very thick

walls and point-like lumina. Often the fibers bear phyto-

liths as spherical stegmata, slightly spinulose, partially

sunken in the basal wall of silica-cell wall. They are visi-

ble even in cross section, and appear in long rows in the

longitudinal sections.

The phytoliths as spherical stegmata also appear, vertical-

ly, on the fibers of fibrous part of the fibrovascular bun-

dles, numerous and in long rows arranged, covering large

surfaces on the fibrous part of the bundle, also visible in

cross-section around caps. Those spherical stegmata seem

to bear rounded spines, and are partially sunken in the

basal wall of the silica-cell. It seems that these stegmata

had from the beginning a definitive size, since it seems

that in time they didn’t grew more. Also, they seem to

connect the fibers with one another, thus enhancing the

solidity of the structure.

Affinities and discussions: In cross section, the studied

material presents a monocotyledonous fascicular struc-

ture, typical for the stem of palm tree, a member of the

Family Arecaceae Berchtold et Presl, 1820 (nom.cons)

also found in the scientific literature as Family Palmae

Jussieu, 1789 (nom. cons. et nom. alt.) or as Family Are-

caceae Schultz-Schultzenstein, 1832 (nom.cons.). The

fossil correspondent, Palmoxylon is a genus name that

generally defines “wood of Palm”, in conformity with the

original diagnosis of Schenk (1882). However, taking

into account that other organs of the palm-plant are con-

nected with the stem (rootlets, leaf axes, petioles, fruits,

flowers, pollen, etc…), there are rare situations, in fossil,

when you can find them together and describe them under

a single name, as in “whole plant” palaeobotanical con-

cept, (see Sakala, 2004), since they fossilize in different

conditions, and usually they are described separately,

under different genus names. Our studied petrified mate-

rial clearly presents fascicular structure so, undoubtedly,

they all can be attributed to the Palmoxylon genus. But

with what kind of extant Palm can be compared? Be-

cause, taking into account the distribution of the anatomi-

cal elements composing the palm-stem we can guess the

position in trunk of the studied sample, aspect which it’s

very important for description, interpretation and identifi-

cation. Anyway the shape, the size, the components of the

fibrovascular bundles and their numeric characters, the

parenchyma of the ground tissue, its abundance and the

arrangement and also the fibrous bundles and the phyto-

liths must be considered in the comparison with extant or

fossil palm structures known from the already published

studies.

The description of the fossil material have followed the

classical model and with language used by botanists and

palaeobotanists which have described the central cylinder

of a palm-stem, in cross section, as formed from three

distinct parts: dermal, subdermal and central zones (see

Mahabalé, 1958 and Kaul, 1960 quoting von Mohl) or

external, intermediary and central in euro-american

school (see Tomlinson, 1961), or subcortical zone (as

periferal sclerotic zone), transitional zone and central

zone as was lastly stated by Thomas & De Franceschi

(2013). From this point of view, the here studied speci-

mens have enough details that send our attention to Cor-

ypha type structure from the subfamily Coryphoideae

Burnett, the tribe Corypheae Martius in Endl.

To do a comparison with extant palms it is difficult since

there are no exhaustive studies done on palm stem anat-

Neogene Palmoxylon from Turkey

37

omy, only partial, due to the difficulties to obtain material

of study, but the studies of Tomlinson, (1961, 1990), and

of Thomas & De Franceschi (2013 with references) must

be considered.

Both the specimens having a splendid fascicular structure

were attributed to Palmoxylon genus. But, again, what

kind of Palmoxylon? Until now over 250 species of

Palmoxylon were described all around the world but few

of them send to an extant correspondent, so it is very dif-

ficult to obtain an answer. However, we tried to find a

similar type of palm, extant or fossil, having the follow-

ing features: arboreal habit, numerous relatively big fi-

brovascular bundles in the central part of the stem, with

sclerenchyma cap of Reniforma type tending to Lunaria

type and fibrous bundles floating in a compact paren-

chymal ground tissue with signs of sustained growing, all

around the fibrovascular bundles only tabular parenchy-

ma in 2-3 regular rows is present. Also, in the fibrovascu-

lar bundles the phloem is usually undivided, rarely appar-

ently divided, and the 2-4(6-8) large metaxylem vessels

have simple or scalariform perforations with thick bars,

and scalariform pitting.

In an important attempt to realize a program of computer-

aided identification for Palm stem anatomy, Thomas

(2011a,b) and Thomas & De Franceschi (2012, 2013)

took the Coryphoideae as case study. In their papers we

found quite interesting suggestions of affinity of our stud-

ied specimens with the typical Coryphoideae. Deeply

comparing the xylotomical characters of our specimens

with the xylotomy of the present day genera as quoted by

Tomlinson (1961, 1990, 2011) and by Thomas & De

Franceschi (2013) we found a similar combination of

features in the genus Corypha L., an extant Palm native

and spread in India, Malaysia, Indonesia, New Guinea,

the Philippines and northeastern Australia (Cape York

Peninsula, Queensland) (Wikipedia, accessed in

20.04.2018). Its species are fan-palms, trees of 20-40 m

high, with the leaves with a long petiole (2-5m) terminat-

ing in a rounded fan of numerous leaflets.

From the fossil forms of Palmoxylon we tried to com-

pared our structures with some fossil forms of so called

"Reniformia group" and we took into account all the

available descriptions of so named "sabaloid Palms" (in

fact from Coryphoideaea subfamily after last phylogenet-

ic classification of Arecaceae of Dransfield et al., 2005),

usually having reniform sclerenchyma caps of the fibro-

vascular bundles, as were described by Schenk (1883),

Berry (1924), Chiarrugi (1933), Rao & Menon (1964),

Menon (1965), Trivedi & Verma (1971a, b), Prakash

(1962), Grambast (1957, 1964), Greguss (1954, 1959,

1969), Prive-Gill & Pelletier (1981), Gottwald (1992),

Nambudiri & Tidwell (1998), Kahlert et al. (2005),

Iamandei & Iamandei (2006), Nour-El-Deen et al. (2017)

where sometimes is specified a possible extant corre-

spondent, being oriented mainly to Sabal, to Trachycar-

pus or to Chamaerops. It was only Grambast (1962) that

has seen in his Palmoxylon sp., of Complanata type, si-

militude with the extant Corypha L.

The resemblance of our structures with the extant Cory-

pha as it is figured and described by Tomlinson (1961),

the similitude shown by our material with the fossil form

described by Grambast (1962) and considered of Corypha

type, confirmed by the papers of Thomas & De Frances-

chi (2012, 2013) support this identification.

The microscopical details observed in our specimens are

similar with those comprised in the species diagnosis

given by Ambwani & Mehrotra (1990) also, regarding the

aspect of the fibrovascular bundles, type of parenchyma

and of stegmata and allow us to attribute the studied ma-

terial to this taxon with the name Palmoxylon coryph-

oides Ambwani & Mehrotra, 1990.

Palmoxylon sp. cf. Trachycarpus

Fig. 6, photos a-i.

Material: HOC25.

Locality: Bolu-Seben-Hoçaş Fossil Site.

Formation: Hançili Formation.

Age: Early Miocene.

Origin on the stem remains: Lower part of an in situ

petrified palm stem.

Storage: This material is now kept under the index

HOC25 within the "Collection of fossil woods" in the

Forest Botany Department, Forestry Faculty, Istanbul

University.

Microscopic description: The studied sample shows in

cross-section only the central zone with fibro-vascular

bundles of collateral type, usually variably oriented, hav-

ing 2-4(-6-8?) metaxylem vessels and not too developed

sclerenchyma caps of Reniforma type to Lunaria type,

floating in the ground parenchymal tissue along with few

small fibrous bundles. Under the sclerenchyma cap a

large median sinus is present, but auricular sinuses were

not observed. The sclerenchyma cap is constituted from

vertically elongated fibrous cells with polygonal rounded

cross-section, thick-walled and with rounded or point-like

lumina. The size of fvb is variable: 650-900/650-750 μm

(mean values: 783/700 µm), the phloematic (=anterior)

sclerenchyma cap has r/tg.d=315/466.6 μm, f/v ratio is

o.68/1 and mean density is of 122.8 bundles/cm2. Signs

of a centrifugal differentiation of the fibrous part seem to

be absent.

Also "the diminutive fibrovascular bundles" (Prasad et al.

2013), seem to be "initials of foliar bundles" and appear

as smaller fibrovascular bundles having well developed

sclerenchyma cap of Reniforma type constituted from

fibers smaller in cross-section and an elongated vascular

part with 2 metaxylem vessels, sometimes tylosed, and

numerous protoxylem vessles, small, grouped. However,

few details can be seen on the phloem and paravascular

parenchyma though they are present in the structure of

the foliar bundle.

The phloem in the normal fibrovascular bundles appear

as protected under the median sinus of the sclerenchyma

cap, in a single island - as an undivided sieve plate and

the phloem cells are discernible.

The metaxylem appears as 2-4 (sometimes 6-8?) vessels,

either oval or elliptic or round (less) in cross-section or,

sometimes, because are partitioned by the very inclined

terminal wall, the apparent number of vessels grows.

They are usually large and thick-walled (9-12 µm the

simple wall), having in cross section 241.6/156.6 μm the

mean diameters. Vertically, the walls of metaxylem ves-

sels have scalariform pitting and thick annular or helical

Stănilă Iamandei, Eugenia Iamandei & Ünal Akkemik

38

Fig. 6 Palmoxylon sp. cf. Trachycarpus H.Wendl. (specimen HOC25). a-c Cross-section. Fibrovascular bundles (fvb)

variably oriented in central zone; compact parenchymal ground tissue, tabular parenchyma around fibros par of fvb; radi-

ant parenchyma around vascular par of fvb; small fvb-s as young foliar bundles; details of fvb-s: metaxylem and proto-

xylem vessels, phloem, reniform sclerenchyma cap, stegmata. d-i Longitudinal sections. Fibrous sheaths of fvb marked

by long rows of stegmata (d). Metaxylem and protoxylem vessels with annular and helical thickenings, scalariform pit-

ting, and with scalariform perforations, parenchymal ground tissue (e-i).

Neogene Palmoxylon from Turkey

39

thickenings. The terminal wall is not very inclined and

bears a scalariform perforation with numerous (5)8-13

thick bars. Often, inside, a fungal attack is present. The

protoxylem is represented by numerous 7-10 (sometimes

more) small polygonal-rounded vessels of 30-80 µm the

diameter and relatively thick walled (4-5 µm the simple

wall). Vertically the protoxylem vessels present thick

annular or helical thickenings sometimes with bifurca-

tions connecting each other, and also, scalariform perfo-

rations.

The intrafascicular or paravascular parenchyma is consti-

tuted of few uniforms, relatively thin-walled (4-6 µm the

double wall) rounded-polyhedral cells which appear po-

lygonal-rounded in cross-section, small-sized cells (d=14-

16 µm), and this because the large vessels occupy the

major part of the vascular part of the fvb. Adjacent to the

metaxylem vessels appear in 1-2 rows regular of flattened

cells, arranged like a tabular parenchyma. Possibly, 1-2

marginal rows of paravascular parenchyma slightly

sclerified cells represent a cup protecting the vascular

zone, forming a small ventral sclerenchyma cap.

The ground tissue or the interfascicular parenchyma is

constituted of slightly emaciated big cells, relatively thin-

walled (3-4 µm the double walls), in fact rounded-

polyhedral cells slightly lobed and very unequal which, in

cross section, appear polygonal-rounded and lobed, of

16/24 - 40/60 µm in diameters. It represents the ground

parenchyma with sustained growing of Corypha type.

Around the sclerenchyma cap of fibrovascular bundles

tabular parenchyma can be seen, in 1-2 compact rows

arranged, but around the vascular part a radiant paren-

chyma appear, sometimes disturbed by compression due

to the sustained growing of the ground parenchyma from

around. The fibrous bundles are absent.

Often the fibers of the sclerenchyma sheath of the fibro-

vascular bundles bear phytoliths as spherical stegmata,

slightly spinulose, partially sunken in the basal wall of

the silica-cell, in long rows arranged, visible in the longi-

tudinal sections. Here also, it seems that these stegmata

connect the fibres one with another, thus enhancing the

solidity of the structure, and probably had not a continu-

ous growing.

Affinities and discussions: The studied specimen has an

obvious monocotyledonous fascicular structure, typical

for a palm-stem, which sustain its attribution to Palmoxy-

lon genus. It shows, in cross-section, characters of the

central zone of Corypha type with fibrovascular bundles

of open collateral type usually variably oriented, includ-

ing also the "diminutive fibrovascular bundles" (see Pra-

sad et al. 2013), possibly as initial of foliar bundles (or

leaf traces) which appear as smaller fibrovascular bundles

having well developed sclerenchyma cap of Reniforma

type, undivided phloem, 2-4(6-8) metaxylem vessels with

annular or helical thickenings, scalariform perforations,

numerous protoxylem vessels, paravascular parenchyma

compact few, tabular-like parenchyma around vessels and

organized as ventral cap, ground tissue as compact paren-

chyma polygonal-rounded and lobed, signs of sustained

growing of Corypha type and also organized around the

fibrous cap of the fibrovascular bundles as tabular paren-

chyma in 1-2 compact rows arranged, around the vascular

part radiant parenchyma appear; fibrous bundles are miss-

ing, phytoliths appear on the fibrous part of the fibrovas-

cular bundles - as spherical stegmata slightly spinulose in

long vertical rows arranged.

Taking into account these details and comparing them

with those comprised in the studies of Tomlinson (1961,

1990) and Thomas & De Franceschi (2013) we believe

that the most similar extant palm is Trachycarpus

H.Wendl., an arboreal palm from the Subfamily Coryph-

oideae Burnett - Tribe Trachycarpeae Satake - Subtribe

Rhapidinae J.Dransf. et al., a palm which is native to

Asia, from the Himalaya east to eastern China. For a spe-

cific affinity of the studied material we took into account

also some published studies on fossil forms with reniform

sclerenchyma cap, or similar.

Thus, Chiarrugi (1933) have described three different

palm-species from the Cretaceous of Somalia: P. bena-

dirense, of Reniformia type, P. scebelianum, of Lunaria

type and P. somalense, of Cordata type. They would be

interesting for comparison but our specimen has very few

details in the vascular zone. Palmoxylon eocenum de-

scribed by Prakash (1962) is also of Reniformia type, but

the problem is the same.

P. parthasarathyi of Rao & Menon (1964) has scleren-

chyma caps of Lunaria and Reniformia type and it was

considered of Cocos type.

Grambast has described in 1957 a Palmoxylon

gignacense of reniform type but not too similar to our

specimen, and in 1962 has described a Palmoxylon sp. of

Complanata type, similar to Corypha, so, different of our

specimen;

Again Grambast (1964) describing a new material bring

into discussion the possible affinities of Palmoxylon

vestitum (Saporta) Stenzel with the extant Phytelephas,

Trachycarpus, Chamaerops.

Other forms of reniform type described by Greguss as

Tertiary "sabaloid palms" from Hungary as Palmoxylon

sabal (?) (Greguss, 1954), P. hungaricum (Greguss,

1959) similar to Livistona and also Palmoxylon doro-

gense, P. sabaloides and P. lacunosum var. axonense

Watelet (Greguss, 1969), which are definitely different of

our studied material.

Trivedi & Verma (1970) have described P. keriense of

reniform-type, considered Cocos-like Palm.

From the 4 forms described by Kramer (1974) only

Palmoxylon sp. form 2 is included into Cordata type, but

the figuration as drawings and photos seem to contradict

the attribution to cordate or lunaria type (sensu Stenzel,

1904), all of them seem to be of reniform type, so it’s

difficult to compare our material with it, but is clearly

different.

Privé-Gill & Pelletier (1981) described a Palmoxylon sp.

of reniform type, however specifying their doubts in a

phrase: “depending of the region of the stem studied, you

can have a lot of groups of Stenzel and Sahni”.

Gottwald (1992) described some specimens of Palmoxy-

lon as P. fasciculosum Vater, 1884 and Palmoxylon sp.

(form 3 - Kramer, 1974) recognizing Lepidocarioid affin-

ities, with the extant Myrialepis and Plectocomiopsis and

respectively with Daemonorops, which could be very

interesting. But Palmoxylon cf. variabile Vater, 1884 has

sabaloid affinities. Nambudiri & Tidwell (1998) have

been described P. hebbertii, also of reniform type, but is

different of our specimen.

Stănilă Iamandei, Eugenia Iamandei & Ünal Akkemik

40

Kahlert et al. (2005) have described a palm of Maastricht-

ian to Palaeocene age from the island Hiddensee (Baltic

Sea) P. bautschii of Vaginata-Reniformia type with nu-

merous metaxylem vessels, but clearly different of our

specimen.

The late Cretaceous species Palmoxylon, techerense de-

scribed by Iamandei & Iamandei (2006), is also of reni-

form type, but has clear Sabaloid affinities.

Other good references were brought by EL-Saadawi et al.

(2004) and Kamal-EL-Din et al. (2013) who described

some fossil palm woods of Egypt, doing also a summary

of the fossil African described Arecaceae ranging from

Cretaceous to Quaternary.

After this discussion on the xylotomic affinities of our

specimen with other fossil forms already described it is

obvious that our specimen has not enough well preserved

details to identify it with a fossil form, or to describe a

new species, even if it is well resembled with the extant

Trachycarpus (see discussion above) so, we attribute it to

Palmoxylon sp. cf. Trachycarpus H.Wendl.

Palmoxylon sp. cf. Borassus L.

Fig. 7, photos a-i.

Material: ERI01.

Locality: Edirne-Erikli.

Formation: volcano-sedimentary.

Age: Middle-Late Miocene.

Origin on the stem remains: Unknown part of the trunk.

Microscopical description: The studied specimen keeps

only the central zone of the central cylinder which is also

of Corypha type (von Mohl, in Thomas & De Franceschi,

2013), with the fibro-vascular bundles sometimes varia-

bly oriented, having very developed reniform scleren-

chyma caps, badly preserved vascular part with a single

one large metaxylem vessel and compact parenchymal

ground tissue. The fibrous bundles are usually missing.

The fibrovascular bundles (fvb) have are very typical

with sclerenchyma cap of Reniforma type, well devel-

oped, with a small median sinus, with rounded auricular

lobes, but without auricular sinuses. The sclerenchyma

cap is constituted from vertically elongated fibrous cells

with polygonal cross-section, thick-walled and with small

rounded or point-like lumina. The size of the fibrovascu-

lar bundles is variable, the mean diameters have

1075/877.7 μm, the phloematic (=anterior) sclerenchyma

cap has the r/tg mean diameters of 836.1/238.9 μm, f/v

ratio is 1.43/1 and the fvb density is of 168.85 bun-

dles/cm2. There are no signs of a centrifugal differentia-

tion of the fibrous part, which is typical for Corypha-type

structures.

The phloem, usually protected under the median sinus,

rarely can be seen as a single island (as undivided sieve

plate) but usually compressed or destroyed by lysis.

The metaxylem appears in cross-section usually as a sin-

gle one round vessel (only sometimes 2-3), often very

badly preserved or not visible. Otherwise all the vascular

part is very badly preserved or is even disappeared. That

round metaxylem vessel seems to be a large one, of about

60-80 μm the diameter in cross section, and is not too

thick-walled. Vertically the walls of metaxylem vessels

have scalariform pitting and annular thickenings slightly

thicker, also difficult to see due to the bad preservation.

Their terminal wall is very inclined and bears a scalari-

form perforation with 7-12 or more thick bars, also only

partially preserved.

The protoxylem, difficult to observe in cross section, is

represented by 3-6 small round to oval vessels. Vertically

the protoxylem vessels present annular thickenings, and

scalariform perforations, details also difficult to observe

due to the bad preservation of the vascular part.

The intrafascicular or paravascular parenchyma cannot be

observed in detail, but seems to be constituted of round-

ed, small uniform, thin-walled cells, rounding and pro-

tecting the xylem vessels.

The ground tissue (the interfascicular parenchyma) ap-

pear in cross section compact as large polygonal rounded

big turgid cells often slightly elongated of 30/50 - 40/70

µm, which are usually thin walled and represents the

ground parenchyma with sustained growing of Corypha

type. Around the fibrous part of the fibrovascular bundles

tabular parenchyma appears, in 1-3 compact rows ar-

ranged, but around the vascular part radiant parenchyma

appears.

Fibrous bundles are usually absent, or very rare.

Phytoliths (as stegmata) on the fibers of the sclerenchyma

caps of the fibrovascular bundles are not obvious, maybe

due to the bad preservation, or simply are missing.

Affinities and discussions: The general aspect of the

structure of the studied sample which is obviously fascic-

ular of palm stem type sustains its attribution to Palmoxy-

lon genus.

The essential xylotomical features of our specimen pre-

serving the central zone of the central cylinder which is

of Corypha type, and has the fibrovascular bundles with

very developed sclerenchyma caps of Reniforma type,

with a small median sinus, with rounded auricular lobes,

but without auricular sinuses, and no signs of a centrifu-

gal differentiation of the fibrous part, which is typical for

Corypha-type structures. Due to the badly preserved vas-

cular part where few details on the phloem - usually

compressed or destroyed by lysis, on the large metaxylem

vessels - one or 2(3?) - which display in longitudinal

view annular thickenings, scalariform pitting, and scalari-

form perforations can be seen, but details of the protoxy-

lem vessels and of the paravascular parenchyma cannot

be observed. The ground tissue appears in cross section

as compact interfascicular parenchyma with sustained

growing of Corypha type. Also, round the fibrous part of

the fibrovascular bundles tabular parenchyma appears, in

1-3 compact rows arranged, and around the vascular part

a radiant parenchyma appears. The fibrous bundles are

usually missing, and phytoliths (as stegmata) on the fi-

bers of the sclerenchyma caps of the fibrovascular bun-

dles were not observed or are missing.

Evaluating all these xylotomical details it seems that our

specimen mostly resembles Borassus L., an arboreal palm

of Subfamily Coryphoideae Burnett - Tribe Borasseae

Mart. in Endl. - Subtribe Lataniinae Meisner, as it ap-

pears described and figured by Tomlinson (1961, 1990),

Dransfield et al. (2008) and Thomas & De Franceschi

(2013). This kind of fan-palm also known as Palmyra

pam, is native to tropical regions of Africa, Asia a New

Neogene Palmoxylon from Turkey

41

Fig. 7 Palmoxylon sp. cf. Borassus L. (ERI01). a-c Cross-section. Central zone with fvb with sclerenchyma cap of Reni-

forma type; compact parenchymal ground tissue; tabular parenchyma around the sclenchyma cap, and radiant parenchy-

ma around the vascular part of the fvb. d-f Longitudinal section. Fibrous sheaths of fvb and compact ground parenchyma.

g-i Metaxylem and protoxylem vessels with annular thickenings, and with scalariform perforations (g).

Stănilă Iamandei, Eugenia Iamandei & Ünal Akkemik

42

Guinea.

For a specific attribution of the studied material we have

taken into account the studies on some published identifi-

cations of fossil forms, considering especially the palms

of Reniforma type. Many forms affiliated to this group

substantially differ of our specimen, sometimes having

affinities with different other extant forms.

Taking into account the similarities with an extant form

of Boraseae, we have chosen a comparison with the few

fossil forms already considered of Borassus type.

So, comparing the structure of our specimen with the

fossil species described by Schenk (in Zittel, 1883),

Palmoxylon zittelii and P. ascheronii we observed a re-

semblance especially with the last one, regarding the

shape of sclerenchyma cap of fvb, and of the parenchy-

mal ground-tissue aspect. Otherwise, later, Palmoxylon

aschersonii Schenck was described again from Paleogene

and Neogene deposits of Algeria and also from the Lower

Miocene of Libya, and was compared with the extant

species Borassus aethiopum Mart. (see Louvet & Mag-

nier, 1971; Boureau, 1947; Boureau et al., 1983)

Petrified Miocene palm stems described from India by

Mahabalé (1959), Sahni (1964), Roy & Ghosh (1980) as

Palmoxylon coronatum have also affinities with the ex-

tant Borassus L. and resembles with our specimen also.

However, since the material don’t offer enough xy-

lotomical details for a specific identification with an al-

ready described form or to describe a new species, we

consider enough to attribute to our here studied specimen

the name Palmoxylon sp. cf. Borassus L.

CONCLUSIONS

Three types of Palmoxylon were identified in this study:

Palmoxylon coryphoides Ambwani et Mehrotra,

Palmoxylon sp. cf. Trachycarpus H. Wendl., and

Palmoxylon sp. cf. Borassus L. This is the first attempt

for description of fossil palms in Turkey, and it is likely

to find new remains of fossil palms especially in the

western Anatolia with further studies.

These three types identified here by xylotomical studies

send to some extant correspondent types. All the locali-

ties of origin of the fossils are situated in a region of Tur-

key where no palm species naturally grows at present

(Fig. 1). The modern representatives of these three types

of palms grow mainly in tropical regions of Africa, east-

ern Asia (including India, China, Malaysia, Indonesia,

New Guinea, the Philippines) and northeastern Australia.

A similar rich palm flora was identified in the Aegean

Greek region (Velitzelos et al. 2017, paper in progress).

The presence of these palm species in the Aegean and

Anatolian regions indicates that the palaeoclimate in the

early Miocene was warmer and wet to subtropical, allow-

ing of the growth of these types of palm trees on the

southern flank of Paratethys Sea. This conclusion regard-

ing the palaeoclimate, at least of the early Miocene, in the

western Turkey also agrees it those given by Akkemik et

al. (2016), Denk et al. (2017 a,b,c) and Güner et al.

(2017).

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